Organization and Formation of the Crossed-Foliated Biomineral Microstructure of Limpet Shells

To construct their shells, molluscs are able to produce a large array of calcified materials including granular, prismatic, lamellar, fibrous, foliated, and plywood-like microstructures. The latter includes an aragonitic (the crossed-lamellar) and a calcitic (the crossed-foliated) variety, whose modes of formation are particularly enigmatic. We studied the crossed-foliated calcitic layers secreted solely by members of the limpet family Patellidae using scanning and transmission electron microscopy and electron backscatter diffraction. From the exterior to the interior, the material becomes progressively organized into commarginal first-order lamellae, with second and third order lamellae dipping in opposite directions in alternating lamellae. At the same time, the crystallographic texture becomes stronger because each set of the first order lamellae develops a particular orientation for the c-axis, while both sets maintain common orientations for one {104} face (parallel to the growth surface) and one a-axis (perpendicular to the planes of the first order lamellae). Each first order lamella shows a progressive migration of its crystallographic axes with growth in order to adapt to the orientation of the set of first order lamellae to which it belongs. To explain the progressive organization of the material, we hypothesize that a secretional zebra pattern, mirrored by the first order lamellae on the shell growth surface, is developed on the shell-secreting mantle surface. Cells belonging to alternating stripes behave differently to determine the growth orientation of the laths composing the first order lamellae. In this way, we provide an explanation as to how plywood-like materials can be fabricated, which is based mainly on the activity of mantle cells.